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The Taxonomic Status of Gladiolus Illyricus (Iridaceae) in Britain

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The Taxonomic Status of Gladiolus Illyricus (Iridaceae) in Britain The Taxonomic Status of Gladiolus illyricus (Iridaceae) in Britain Aeron Buchanan Supervisor: Fred Rumsey, Natural History Museum, London A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science of Imperial College, London Abstract First noticed officially in Britain in 1855, Gladiolus illyricus (Koch) presents an interesting taxonomic and biogeographical challenge: whether or not this isolated northern population should be recognized as a separate sub-species. Fundamental conservation issues rest on the outcome. Here, the investigation into the relationship of the G. illyricus plants of the New Forest, Hampshire, to Gladiolus species across Europe, northern Africa and the middle east is initiated. Two chloroplast regions, one in trnL–trnF and the other across psbA–trnH have been sequenced for 42 speci- mens of G. illyricus, G. communis, G. italicus, G. atroviolaceus, G. triphyllos and G. anatolicus. Phylogenetic and biogeographical treatments support the notion of an east–west genetic gradation along the Mediterranean. Iberia particularly appears as a zone of high hybridization potential and the source of the New Forest population. Alignment with sequences obtained from GenBank give strong support to the classic taxonomy of Gladiolus being monophyletic in its sub-family, Ixioideae. Comments on these chloroplast regions for barcoding are also given. In conclusion, the genetic localization of Britain’s G. illyricus population as an extremity haplotype suggests that it could well deserve sub-species status. Contents 1 Introduction 2 2 Background 4 3 Materials and Methods 8 4 Results and Discussion 15 5 Conclusions 26 Appendices 28 References 56 1. Introduction G. illyricus in Britain Figure 1: G. illyricus flower spike. Photograph: Fred Rumsey, Denny Wood, New Forest. 1 Introduction The study of Gladiolus illyricus (Koch) is particularly pertinent at this time. It is a medium-sized flowering monocotyledon with a Mediterranean distribution, but with one important exception: the small yet significant, and now declining population in the New Forest of Hampshire, here in the British Isles. The natural history of G. illyricus in the UK, until as recently as the 1950s, ranges from being unclear to unknown. The first recorded sighting was on the Isle of Wight in 1855, with notes of its presence in the New Forest appearing soon thereafter. The origins of this population, in the extreme north of the species’ range, immediately became a point of debate. Was the arrival part of a natural progression northwards or was it an artificial, human-mediated introduction? Today, the continuing survival of G. illyricus in the New Forest appears fragile, mak- ing the above an important question. It is currently a protected Schedule 8 species and so protected under law by the Wildlife and Countryside Act, 1981, but with its endemic status unresolved, it remains on the waiting list of Cheffings and Farrel (2005)∗. The wait- ing list is populated by “taxa for which questions still remain over taxonomic validity or endemic status.” Indeed, the example of such a taxon given by the authors is G. illyricus ssp britannicus. This state of conservation limbo is unsatisfactory, for if the endemic sta- ∗‘The Vascular Plant Red Data List for Great Britain’ is part of the Joint Nature Conservation Com- mittee’s Species Status project, which is used to inform the statutory five-yearly review of Schedules 8 species. 2 1. Introduction G. illyricus in Britain tus is rejected then much demanded conservation resources can be redirected elsewhere. However, if it is confirmed, a fuller more concerted effort would be justified. The issue of the British nativity of G. illyricus is theoretically inseparable from the suggestion that the population in the New Forest should be designated as being a separate subspecies, G. illyricus ssp britannicus. This study was born with the desire to resolve the taxonomic status of G. illyricus and hence shed light on the natural history of the New Forest population. Resolution would thus make clear the level of impetus required of conservation efforts. An outline for a study to answer the question has been proposed by Lockton (2006); that is, to demonstrate: 1. the existence of an east–west genetic gradation 2. that the gradation is correlated to natural dispersal rates. 3. that there is speciation within that gradation. 4. that the British plants are sufficiently distinct. 5. that they belong to a distinctive semi-natural vegetation habitat. The ecological investigation, that can make clear the issues of the fifth point has al- ready started, most notably by Stokes (1987). This study begins the molecular investiga- tion of the G. illyricus population of the New Forest, and hence starts to address the issues of the first, third and fourth points. The work began with the important grounding step of placing the European Gladiolus species into a wider context. Gladiolus is one of the largest genera in the Iridaceae family, with possibly as many as 300 species. Almost all of these grow in sub-Saharan Africa (Goldblatt et al. 2001); less than twenty species are endemic to Europe and neighbouring countries. Support for the monophyletic status of all Gladiolus is needed to be able to rely on existing work that assumes the long standing taxonomy. After the broad initial analysis, the project then focussed on the inter-relations within Europe, which were then explored with the above framework in mind. Sadly, the full answer can not be given here: limited time restricted the scope of what was achievable. However, important insights into the wider situation are revealed, providing some answers and raising significant new ones, thus informing future research. 3 2. Background G. illyricus in Britain Figure 2: An example of G. illyricus from the herbarium at Reading University. This is a New Forest plant, collected in 1960. 2 Background The first systematic study of Gladiolus illyricus (Koch) growing in the UK did not start until almost one hundred years after it was first officially noticed. As such, our knowledge of its distribution and ecology is very sparse for times before 1950, when Bowman started the surveying of the New Forest for this purpose (Rand, 2005), and almost completely blank for anything before 1855, when Mrs Phillipps of the Isle of Wight made the first record of its growing in the British Isles (More, 1862). According to Townsend (1904) the first mainland discovery was in 1856 by Rev. W. H. Lucas. From the mid to the end of the 19th century, the story seems to be one of undirected specimen collection with the occasional serendipitous discovery of new sites. The Natu- ral History Museum’s British Herbarium holds thirty nine specimens from this period (the first being the 1858 sheets of J. T. Boswell-Syme), all collected from within the New For- est (v.c.11), seldom with a more detailed description of location than “near Lyndhurst”. Accompanying notes are generally sparse as well, although Rand (2005) points out that descriptions of habitat, beyond “amongst bracken”, were sometimes made. The most detailed of these are by Dyer and Trimen (1864), but they worked on only two sites in the New Forest, within less than 3km of each other. Later, Townsend (1904) compiled a summary of the New Forest locations where it had been found, describing three almost distinct areas of modest size, all within 6km of Lyndhurst. Crucially, Pope et al. (2005) 4 2. Background G. illyricus in Britain notes that it was also found near Ensbury, Dorset (v.c.9) around 1874 and collected trice more from the Isle of Wight in 1872, 1897 and 1931, although the last collection was rather haphazard. The plants seems to have then received little attention from the turn of the century until the systematic surveys of Bowman from the 1950s onwards, Hamilton in the 1960s and Everett in the 1980s (Rand, 2005), as well as Stokes (1987). From their foundational reports and the continued efforts of the New Forest Study Group, we are now fairly certain of the full extent of the sub-populations growing in the New Forest. The widespread activity of botanists across Britain means that we can now be sure that G. illyricus no longer grows outside this area. The current range, while still small, is sev- eral times larger than that investigated by botanists at the turn of the century, spreading up to 12km from Lyndhurst. What the actual distribution was in those early years is almost impossible to deduce, but it seems reasonable to surmise, with sub-populations growing outside the New Forest at the time, that it was more consistently widespread then, than it is today. However, this is a pivotal historic factor for the question of G. illyricus being endemic in the New Forest: care must be taken before assumptions are made. Was the sudden start of botanical inter- est in the middle of the 1800s perhaps a result of a contemporary introduction to the south coast from a continental population? I believe not: it is most likely due to artifacts of historical circumstances and not because of any events or trends belonging to the plants themselves over the period in question. Indeed, Mrs Phillipps did not luckily stumble upon the singular occurrence of G. illyricus on the Isle of Wight, but rather that the “Wild Gladiolus” plant had been growing in the woods at Shanklin, the nearby town, for at least a generation (More, 1862). It is quite possible that, had local people written of the flow- ers that they commonly saw, or such notes more widely disseminated, records of many plants would have appeared much earlier. Martin Rand (2005) makes a good case for the existence of G. illyricus in the New Forest before the first records appeared.
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